Studies of Malagasy Eugenia – IV: Seventeen new endemic species, a new combination, and three lectotypifications; with comments on distribution, ecological and evolutionary patterns

Abstract Seventeen new endemic species of the genus Eugenia L. (Myrtaceae) are proposed from Madagascar, including: Eugenia andapae N. Snow, Eugenia barriei N. Snow, Eugenia bemangidiensis N. Snow, Eugenia calciscopulorum N. Snow, Eugenia delicatissima N. Snow, Callm. & Phillipson, Eugenia echinulata N. Snow, Eugenia gandhii N. Snow, Eugenia hazonjia N. Snow, Eugenia iantarensis N. Snow, Eugenia malcomberi N. Snow, Eugenia manomboensis N. Snow, Eugenia obovatifolia N. Snow, Eugenia ranomafana N. Snow & D. Turk, Eugenia ravelonarivoi N. Snow & Callm., Eugenia razakamalalae N. Snow & Callm., Eugenia tiampoka N. Snow & Callm., and Eugenia wilsoniana N. Snow, and one new combination, Eugenia richardii (Blume) N. Snow, Callm. & Phillipson is provided. Detailed descriptions, information on distribution and ecology, distribution maps, vernacular names (where known), digital images of types, comparisons to morphologically similar species. Preliminary assessment of IUCN risk of extinction and conservation recommendations are provided, including Vulnerable (4 species), Endangered (2 species), and Critically Endangered (4 species). Lectotpyes are designated for Eugenia hovarum H. Perrier, Eugenia nompa H. Perrier, and Eugenia scottii H. Perrier respectively.


Introduction
Th is paper is the fourth in a series devoted to clarifying the systematics of Malagasy Eugenia L. (Snow 2008(Snow , 2011Snow et al. 2012), which until the work of Miller (2000) and Labat and Schatz (2002) had been dormant for fi ve decades since early treatments by Perrier de la Bâthie (1953a,b). Its purpose is to propose seventeen new species, make a new combination, and lectotypify three names. It also provides detailed species descriptions with comparisons to morphologically similar taxa, distribution maps, vernacular names, digital images of types, and preliminary conservation assessments following IUCN Red List Categories and Criteria (2012).

Materials and methods
Type specimens of all nearly described species of Eugenia from Madagascar have been examined. All Malagasy specimens of Eugenia housed at MO (abbreviations following Th iers (2015)) have been viewed by NS (through May 2014), as have all duplicates housed at his institution (KSP). Specimens housed at G and P have been studied by MC and PP. Duplicates of many specimens housed at KSP also have been consulted by NS. Specimens cited here also were compared against all material of Eugenia from the Comoros and Mascarenes housed at MO through May 2014. All specimens housed at MO should have duplicates at P and TAN given existing exchange agreements, although none at TAN have been seen by the authors. Accession numbers of specimens (where indicated) follow the herbarium acronym with a hyphen, whereas barcode numbers are enclosed in square brackets. (15-)18.5-25 × (7-)13.5-14 cm, broadly elliptic to broadly obovate, base cordate and somewhat clasping (or rounded), surface and margin fl at, apex obtuse to broadly acute, tip acuminate (or rarely retuse), secondary veins more or less straight, 11-15 per side protruding prominently below and arising at 25-55° angles and connected by slightly arching (but also prominent) inner marginal veins, tertiary veins relatively well-spaced but projecting only slightly (dried material); adaxial surface glabrous, oil glands faint (use magnifi cation), sparse to common, more or less fl ush and darkish (dried), midvein sulcate; abaxial surface glabrous, midvein projecting prominently throughout and punctate (especially proximally) or epunctate, secondary veins projecting prominently, straight or only curving slightly towards margin, the secondaries connected at their ends by moderately arching connecting veins, tertiary veins projecting but less so than secondaries, intramarginal vein of same thickness as tertiaries, 1.5-5 mm from margin at midpoint of blade. Infl orescence (material scant) a monad; fl owers caulifl orous, arising from short brachyblasts (< 3 mm long) above nodes on naked branches. Pedicels 8-32 mm long (possibly elongating after fertilization), 0.7-2.0 mm wide, somewhat compressed laterally, longitudinally striate, somewhat fl exuous (bending with light touch), habit unknown, moderately glandular (glands faint), anthopodium and metaxyphylls absent. Bracteoles narrowly to broadly ovate, 1.5-2.5 × 0.5-1 mm, minutely and sparsely hairy dorsally and apically (hairs clear or whitish with some reddish). Hypanthium campanulate, 3.0-3.3 mm long, 3-4 mm wide at base of calyx lobes, densely but very shortly sericeous in proximal half (hairs reddish-brown) but glabrous distally; ovary apex glabrous. Calyx lobes 4 and often tearing irregularly towards hypanthium, up to 3.5 mm long × 4.5 mm broad (at base), irregularly hemispherical, glabrous on both faces apart from occasional minute hairs, evidently refl exed irregularly in anthesis. Petals 4 (material scant), 5.5-19.0 × ca. 10 mm, narrowly to broadly obovate, glabrous, epunctate, rose to violet. Staminal ring 3.5-4.0 mm in diameter (rounded or somewhat squarish), sparsely short hairy (hairs whitish). Stamens ca. 140 (estimated from scars on ring), multiseriate; fi laments up to 10 mm long; anthers globular, ca. 0.8 mm long. Fruit 23-35 × 21-50 mm, depressed globular to globose, glabrous, base and apex rounded or apex crowned by calyx lobes, pinkish-reddish.
Phenology. Flowering in February and March; fruiting March through November. Etymology. Th e species occurs in the Andapa Basin. Distribution. Known in northeast Madagascar in and around the Anjarahabe-Sud and Marojejy protected areas (Figure 7).
Habitat and ecology. Humid forests, riparian areas and near summit of Ambodisatrana;ca. 200-1540 m. Conservation status. With an Extent of Occurrence (EOO) of 1586 km 2 , and Area of Occupancy (AOO) of 45 km 2 and fi ve subpopulations, two of which are situated within the protected area network (Anjanaharibe-Sud, Marojejy), Eugenia andapae is assigned a preliminary risk of extinction of "Vulnerable" [VU B1ab(iii)+2ab(iii)] following the IUCN Red List Categories and Criteria (IUCN 2012). Comments. Th e calyx lobes of this species tear towards the hypanthium during anthesis (see also E. lacerosepala N. Snow and E. ambanizanensis N. Snow). In this regard Eugenia andapae is similar to taxa fi rst described by Scott (1979) from the Mascarenes in Monimiastrum A.J. Scott, which were reduced to synonymy under Eugenia (Snow 2008). Th e relatively long and broadly elliptic to obovate leaf blades with cordate bases, and the relatively straight and projecting abaxial secondary veins are diagnostic among other species of Eugenia in Madagascar. On herbarium material the abaxial tertiary veins also protrude slightly despite being thin. Th e label of the type specimen indicates that the fl eshy fruits typically are crowned by the calyx lobes, although this was not true for the specimen from the Beamalona River. Description. Shrubs to 3 m tall; bark of main bole unknown. Vegetative and reproductive parts (where indicated) bearing a moderately dense, shortish indumentum, the individual trichomes dibrachiate or not, appressed to somewhat refl exed (appearing villous), frequently irregularly contorted, whitish or reddish. Branchlets laterally compressed but becoming rounded, smooth, moderately short villous (hairs mostly refl exed and not dibrachiate) becoming glabrous, oil glands common and prominent (after indumentum falls away). Leaves opposite, mostly occurring in 2-4 pairs along seasonal growth of branchlet, thinly coriaceous, venation brochidodromous (invisible to obscure), discolorous, somewhat glossy above but matte below. Axillary colleters absent. Petioles 1.7-2.5 mm long, broadly sulcate above, moderately hairy towards base adaxially in sulcus. Leaf blades 0.9-2.0 (-2.7) × 0.6-0.9 cm, narrowly elliptic or elliptic to narrowly obovate, base cuneate, surface fl at to slightly and irregularly (but broadly) sinuous on drying, margin fl at or drying slightly revolute, apex obtuse; abaxial surface glabrescent, oil glands common (use magnifi cation) and drying brownish and slightly sunken, midvein fl ush and becoming imperceptible towards apex; abaxial surface sparsely glabrescent, oil glands relatively sparse to moderate and somewhat less prominent than adaxially, secondary veins few and barely perceptible, the secondaries connected at their ends by a slightly arching pseudo-intramarginal vein 0.3-0.8 mm from leaf margin (i.e., lacking an intramarginal vein distinct from pseudo-intramarginal vein). Infl orescence a monad; the base of the fl owering branchlets each with (2-)4-6 fl owers arising alternately, each fl ower subtended by a short, hairy and somewhat ovate to broadly triangular caducous bract. Pedicels (5-)10-15(-20) mm long, 0.3-0.5 mm wide, round in transverse section, stiff , ascending, sparsely hairy (especially near base) to nearly glabrous, moderately glandular throughout, anthopodium present or absent. Bracteoles 2, linear, 1.0-1.2 × 0.3-0.5 mm, sparsely hairy. Hypanthium cupulate 2.0-2.5 mm long, 1.4-1.8 mm wide at base of calyx lobes, densely short-hairy, oil glands absent or sparse (and obscured by hairs); ovary apex glabrous. Calyx lobes 4, 1.5-1.9 mm, broadly ovate to rounded, glabrous on both faces apart from sparse apical hairs (white or reddish), strongly refl exed in athesis. Petals 4 (material scant), ca. 2.5 mm × 2 mm, obovate to widely obovate, glabrous on both faces apart from sparse apical hairs (contorted irregularly), oil glands absent. Staminal region (i.e., lacking a well-defi ned staminal ring) 1.6-1.8 mm diameter in anthesis, sparsely hairy (trichomes simple); stamens 35-45; fi laments 1.5-2.5 mm; anther sacs 0.5-0.7 mm long, globose, basifi xed, eglandular. Style 2.5-2.8 mm, glabrous or sparsely hairy basally; stigma narrow and only slightly capitate. Fruit unknown.
Etymology. Th e specifi c epithet honors Dr. Fred Barrie (b. 1948) of the Missouri Botanical Garden in recognition of his contributions to our knowledge of Eugenia and other genera of Mesoamerican Myrtaceae (e.g., Barrie 2004Barrie , 2005.
Phenology. Flowering confi rmed only for the middle of November; fruiting likely late November through December.
Distribution. Known only from near Port Bergé in Mahajanga Province ( Figure 4). Conservation status. With only one collection known from Central-western Madagascar collected in an unprotected and threathened dry forest, Eugenia barriei is assigned a preliminary risk of extinction of "Critically Endangered" [CR A3c] following the IUCN Red List Categories and Criteria (IUCN 2012). In the absence of eff ective protection and the high human pressure on these forests, it is unlikely that the forest will persist beyond 3 generations of Eugenia barriei (ca. 30 years).
Comments. Th e type specimen of Eugenia barriei initially was determined as E. tropophylla H. Perrier. Th e latter species and the varieties described by Perrier de la Bâthie (1953a,b) do not represent a single taxon, which even a cursory glance at the numerous syntypes (at P) will reveal. Among taxa from southeastern Africa, Eugenia barriei resembles some specimens of E. capensis subsp. gracilipes. In particular, the slender pedicels of a specimen from Malawi (Chapman 6570 [MO]) have a similar but less dense indumentum on the branchlets, pedicels and hypanthium. Other diff erences of the Chapman specimen include longer and more densely and prominently punctate leaves, and infl orescences that mostly arise from ramifl orous brachyblasts.
Etymology. Th e epithet is derived from the place name Bemangidy, a biologically rich region within the provisional Protected Area of Tsitongambarika.
Phenology. Flowering commencing November and December; fruiting likely commencing by late December or early January (unconfi rmed).
Habitat and ecology. Label information is sparse, but based on recent satellite imagery, evidently occurring in moderately to highly disturbed hilly areas that retain thin remnants of forest in ssome drainages and on steeper slopes, the type gathering located some 0.5 km east of much less disturbed primary forests that occur on steeper slopes, the paratype gathering in highly disturbed sites; elevation ca. 25-110 m.
Conservation status. With only three collections known, an AOO of 9 km 2 and one subpopulation, which is situated outside the current protected area network, Eugenia bemangidiensis is assigned a preliminary risk of extinction of "Critically Endangered" [CR A3c+B1ab(iii)] following the IUCN Red List Categories and Criteria (IUCN 2012). All the known material of the species was collected in the proposed new protected area of Tsitonggambarika in the south-east (Fig. 2). Th e new species seems to have a highly restricted distribution and the lowland evergreen tropical forests where the species grows is under threat. Its defi nitive protection would likely allow this species to be downlisted to "Endangered".
Comments. Th e leaf morphology of Eugenia bemangidiensis resembles E. wilsoniana (see discussion above). However, the short-pedicellate, fasciculate, and caulifl orous infl oresences of E. bemangidiensis diff er from the axillary, long-pedicellate fl owers of E. wilsoniana. Th e type gathering at ca. 25 m elevation of E. bemangidiensis is approximately 645 km south of the most southerly collections of E. wilsoniana, the latter of which occur at middle elevations of ca. 980-1100 meters.
Etymology. Calci and scopulorum combine to form a genitive plural masculine noun, in reference to the locality at the base of a calcareous (limestone) cliff .
Distribution. Known only from the type gathering in north-eastern Madagascar in Antsiranana, in the Montagne des Français limestone massif (Figure 7).  Habitat and ecology. Degraded dry forest at the foot of a cliff over calcareous substrates; ca. 410 m.
Conservation status. Th e species was said to be abundant at the type gathering. However, with only one collection known, an AOO of 9 km 2 and one subpopulation, which is situated within a proposed protected area that currently holds a temporary protection status (Montagne des Français), Eugenia calciscopulorum is assigned a preliminary risk of extinction of "Critically Endangered" [CR A3c+B1ab(iii)] following the IUCN Red List Categories and Criteria (IUCN 2012). Th e species seems to have a highly restricted distribution and the dry deciduous forests around Montagne des Français is mostly degraded as a result of human activities and is under serious threat. Defi nitive protection of those dry forests likely would allow the species to be downlisted to "Vulnerable".
Etymology. A superlative of delicata in Latin, in reference to the thin, almost thread-like pedicels. Vernancular name. Rotramadinika (Rakoto 294). Phenology. Flowering in late October through at least early November; fruiting November.
Distribution. Known from the northern mountains of Madagascar in and around the Manongarivo and Tsaratanana protected areas and in the Sorata region (Fig. 4).
Habitat and ecology. High elevation montane forests with bamboo; 1100-2350 m. Conservation status. With an EOO of 3278 km 2 , an AOO of 36 km 2 and four subpopulations, two of which are situated within the protected area network (Manongarivo, Tsaratanana), Eugenia delicatissima is assigned a preliminary risk of extinction of "Vulnerable" [VU B1ab(iii)+2ab(iii)] following the IUCN Red List Categories and Criteria (IUCN 2012). At the summit of Beampoko Eugenia delicatissima was indicated as being rare (Rakotovao 2565).
Comments. Eugenia delicatissima is part of an apparent species group characterized by thinly coriaceous, sinuous leaf blades and slender, often elongate and delicate pedicels.
Th e thinly coriaceous, sinuous leaf blades of Eugenia delicatissima are somewhat suggestive of E. echinulata, but the fruits of E. delicatissima are smooth (e.g. Randriambololomamonjy 270), unlike the highly irregular (warty to echinate) texture of of E. echinulata.
Th e slender pedicels are similar to those of Eugenia tropophylla H. Perrier, a highly heterogenous species that Perrier de la Bâthie (1953a: 167) initially described with three subspecies, none of which match E. delicatissima. Th e taxonomy of E. tropophylla needs serious revision and will be the subject of a forthcoming publication.
Etymology. Th e specifi c epithet comes from the Latin echinulatus (having very small prickles) in reference to the prickly, dried, mature fruits that are unique for the genus in Madagascar.
Vernacular name. Rotra (Antilahimena 6841). Phenology. Flowering December; fruiting January and February. Distribution. Known from the eastern escarpment of Madagascar in the Toamasina Province around Analamazaotra and Ambatovy and in the highlands at Anjozorobe in Antananarivo Province (Fig. 7).
Habitat and ecology. Dense humid evergreen forests; ca. 915-1350 m. Conservation status. One collection indicated the species as being abundant in its area of collection (Randrianaivo 1463 et al.). However, with an EOO of 4,372 km 2 , an AOO of 72 km 2 and three subpopulations, two of which are situated within the protected area network (Analamazaotra, Zahamena), Eugenia echinulata is assigned a preliminary risk of extinction of "Vulnerable" [VU B1ab(iii)+2ab ( Comments. Eugenia echinulata is unknown in fl ower apart from one remnant hypanthium on the holotype. However, the coarsely sinuate leaf blades and acuminate apices, coupled with the warty mature fruits, are diagnostic and unknown in combination elsewhere among Malagasy congeners. Whereas E. muscicola H. Perrier has narrowly elliptic leaves, its leaf margins are more or less planar. A number of similar sterile collections from this area may be this taxon. Th e texture of the fruit is coarsely rugulose-warty (see link above to image of living material), but the warty protrusions collapse to a large extent and upon drying and then assume a sharper, echinulate texture. Description. Shrubs to trees, 3-12 m. Trunk 3-8 cm dbh; outer bark of main bole reddish, papery to fl akey and peeling. Foliage and reproductive parts glabrous except where noted. Branchlets terete to laterally compressed, reddish when fresh but drying brownish or grayish, glabrous but prominently glandular, the epidermis smooth but soon peeling (dried specimens) and often thin and narrow grayish strips or even threads. Leaves opposite, medium green (fresh), slightly to strongly discolorous, venation brochidodromous, thinly coriaceous (easily cracking when dried), surfaces matte. Axillary colleters lacking. Petioles 2.8-6 mm, slightly sulcate distally. Leaf blades (5.5-)9-16 × 2.5-6.0 cm, elliptic to occasionally obovate, base cuneate, apex acute to mostly acuminate or caudate, margins fl at; adaxial surface glabrous, densely punctate (glands small and best seen with magnifi cation); abaxial surface similar, lateral veins indistinct to prominent; intramarginal vein 1-3.5 mm from margin at midpoint of blade. Infl orescence terminal, axillary, or ramifl orous, of soliltary monads or in fascicles of 2-3 fl owers. Pedicels 9-29 mm long × 0.8-1.2 mm thick, stiff , moderately glandu- lar, ascending to erect, green. Bracteoles narrowly ovate to broadly rounded, 0.5-1.3 × 1.0-1.2 mm, stiff but thin, glabrous to minutely and sparsely sericeous on margin, often persistent in fruit. Hypanthium 4-5 mm long, cupuliform, densely glandular (some glands larger proximally). Calyx lobes 4, 6-9 × up to 11 mm, broadly elliptic to rounded, cream-colored (fresh), refl exed in fl ower, persistent and often crowning the fruit. Petals 4, 11-14 × up to 12 mm, widely elliptic to widely obovate (narrowly greatly at base), sparsely and minute ciliate marginally, densely glandular (glands of variable size but typically large proximally). Stamens 250-300, multiseriate, exserted; staminal disk short-hairy (trichomes slightly ferrugineous); fi laments 8-15 mm, white; anthers globose, 0.3-0.5 mm, brownish, bearing a single large apical gland. Ovary apex glabrous but somewhat glandular. Style 8-10 mm; stigma narrow (scarcely if at all swollen). Berries 2.5-4.0 × 3.0-4.0 cm, subglobose to globose, purplish or violet (drying nearly black). Locules 2; placentation axile. Seeds up to 20 mm long and wide (available material possibly not fully mature), up to 6 per fruit, evidently often somewhat fl attened, cotyledons and hypocotyl barely if at all diff erentiated.
Etymology. Th e species honors Dr. Kanchi N. Gandhi (b. 1948) of Harvard University Herbaria, North American editor of the International Plants Names Index, and an expert of botanical nomenclature who has assisted colleagues with the proper usage of botanical Latin and interpretation of the Code over many years. Vernacular names. Gavoala (Ravelonarivo 103); gavoalabe. Th e name "guaavaala" appears on one specimen, but native speakers inform us that it is a nonsensical name based on a combination of the common English name guava (for members of Psidium, a Neotropical genus) and ala, meaning locally "of the forest".
Phenology. Flowering January through June; fruiting March through November. Distribution. Endemic to northeast Madagascar, recorded from the Island of Nosy Mangabe, the Masoala Peninsula, and in and around the Anjaniharibe Reserve ( Figure 11).
Habitat and ecology. Humid and typically dense forests; 0-1424 meters. Conservation status. With an "Extent Of Occurrence" (EOO) of 9,504 km 2 , an "Area Of Occupancy" (AOO) of 45 km 2 and fi ve subpopulations, three of which are situated within the protected area network (Anjanaharibe-Sud, Marojejy and Nosy Mangabe), Eugenia gandhii is assigned a preliminary status of "Least Concern" [LC] following the IUCN Red List Categories and Criteria (IUCN 2012).
Comments. Perrier de la Bâthie's E. arthroopoda H. Perrier var. ambalavensis H. Perrier was based on a single collection from the lowlands just to the south of Marojejy Reserve, but we have little doubt that it is conspecifi c with the material we include in E. gandhii, which ranges from the nearby Anjaniharibe Reserve to the southwest of Marojejy to the Masoala peninsula and Nosy Mangabe farther south.
Etymology. Named after the local vernacular name hazonjia, which is recorded on two specimen labels.
Phenology. Flowering September through December; fruiting unknown. Distribution. Known from Antsiranana Province in north-eastern Madagascar approximately 30 km northwest of Vohemar and in the Daraina region (Fig. 11).
Habitat and ecology. From littoral forest on sand and dry deciduous forest; 15-1015 m.
Conservation status. With an EOO of 39 km 2 , an AOO of 27 km 2 and two subpopulations, one of which is located within in the temporarily protected Loky-Manambato area, Eugenia hazonjia is assigned a preliminary risk of extinction of "Endangered" [EN B1ab(iii)+2ab(iii)] following the IUCN Red List Categories and Criteria (IUCN 2012). Th e new species appears to be hightly restricted in its distribution, and the lowland evergreen tropical and littoral forests where it grows is under threat. Its defi nitive protection likely would allow downlisting to "Vulnerable".
Etymology. Th e specifi c epithet is a Latinization of the Iantara River, location of the type gathering.
Phenology. Flowering March through November; fruiting November and December. Distribution. Known from the eastern escarpment of Madagascar in Finarantsoa and Toamasina provinces around and in the Andringitra, Ranomafana and Analamazaotra Special Reserve (Fig. 11).
Habitat and ecology. Evidently along or near rivers in humid forest, sometimes in disturbed areas; from 770-1210 m. Th e type collection indicates the plant, presumably the large fruits, is used by the Common brown lemur, Eulemur fulvus.
Conservation status. With an EOO of 4,892 km 2 , an AOO of 81 km 2 and three subpopulations, all of which are situated within the protected area network (Andringitra, Ranomafana and Analamazaotra), Eugenia iantarensis is assigned a preliminary status of "Least Concern" [LC] following the IUCN Red List Categories and Criteria (IUCN 2012).
Comments. Th e fl owering and early fruiting material (Toamasina Province) and mature fruiting material (Finarantsoa) are disjunct and the fl owering material is somewhat incomplete (no petals), but apart from slightly shorter leaves in Toamasina the specimens match well.
Mature fruits are among the largest yet known among Malagasy Eugenia, and the large number of seeds per fruit (up to 13) is considerably greater than that commonly seen for Eugenia worldwide or in Madagascar. Th e testa in dried material becomes highly furrowed-convoluted to resemble the surface of a human brain, likely due only in part to shrinkage from dessication. A similar testal morphology was recently described in Eugenia alletiana Baider & Florens from Mauritius (Baider and Florens 2013), although the testa of that species at maturity was said to be as fi rm as the texture of wood. Th e selective pressures underlying the evolution of such a thick and tough testa are worthy of consideration in light of the observation on the holotype label that the trees are used by brown lemurs. Th e walls of the fruit and testa frequently have bore holes of an unknown insect of ca. 2 mm diameter.  Description. Trees to 6 meters tall; bark of main bole unknown. Indumentum (where noted) of vegetative and reproductive material sericeous (hairs dibrachiate and typically reddish or brownish). Emerging (youngest) branchlets laterally compressed, sparsely hairy, glandular, color uncertain; older branchlets becoming terete, soon glabrous, mostly eglandular, becoming brownish-gray, smooth. Leaves evenly distributed along branchlets (nodes greater than 9 cm apart), coriaceous, venation brochidodromous, strongly discolorous, matte above and below. Axillary colleters present (type A; Figure 14. Holotype specimen of Eugenia malcomberi (MO). Snow et al. 2003) on youngest emerging leaves, otherwise absent. Petioles 8-11 mm long, fl attened above, sparsely hairy upon emergence but becoming glabrous, glandular (particularly above adjacent to blade). Leaf blades 14.7-22.6 × (9.3-)10.0-)12.5 cm, broadly elliptic (mostly) or ovate, base rounded, margin and blade surface fl at, apex and tip acute; abaxial surface sparsely hairy at emergence becoming glabrous, oil glands prominent (becoming less so), midvein broad (1.5-2.0 mm wide) and fl ush at emergence but becoming slightly sulcate in older leaves at least proximally; abaxial surface glabrous, oil glands sparse to moderate on laminar surface but common on midvein, midvein protruding but becoming fl ush or nearly so towards apex, secondary and intramarginal veins prominent, the veins connecting secondaries also prominent and broadly arching; intramarginal vein relatively faint, irregularly sinuous and 2-3 mm from midpoint at laminar margin. Infl orescence mostly ramifl orous or axillary, fasciculate clusters and comprised monads and triads to 5-7-fl owered cymes. Bracteoles 1.1-1.8 mm × ca. 0.5 mm, ovate, sparsely hairy, glandular, mostly persisting during anthesis. Hypanthium 4.3-4.5 × 4.0-5.3 mm wide below base of calyx lobes, cupulate, sparsely to moderately hairy especially towards base, oil glands common to dense. Calyx lobes 4, 3.5-6.5 mm long, rounded to broadly obovate, apex obtuse, sparsely ciliate, oil glands moderately common and projecting (especially lower surface), consisting of two longer (inner) lobes and two shorter outer lobes (the bases of the outer lobes covering the bases of the inner lobes in bud), evidently whitish in fl ower. Petals 4 (5 on one fl ower), 11-17.5 × 6-12 mm, obovate to widely obovate, glabrous, white, oil glands sparse to common and pronounced. Staminal ring ca. 3.5-4.5 mm in diameter, glabrous. Stamens (estimated) 100-150; fi laments 5-13 mm long; anther sacs 0.7-0.9 mm long, globose to subcylindrical; connective bearing one faint apical gland or eglandular. Ovary apex glabrous. Style 10-13 mm long, glabrous, eglandular; stigma narrow (barely if at all swollen). Berries not seen, indicated as being green on specimen label.
Epynomy. Th e new species honors Dr. Simon Malcomber (b. 1967), who collected the type material and nearly two thousand specimens from Madagascar in the early 1990s.
Phenology. Flowering in early February; fruiting probably late February through at least mid-March.
Distribution. Known only from the type gathering in Toliara in parcelle I of the Reserve Naturelle Integrale #11 northwest of Taolognaro, in south-western Madagascar. Th is region is near the southern terminus of the humid forests that occur east of the main escarpment that run latitudinally along much of the length of Madagascar (Fig. 4).
Habitat and ecology. Th e species was collected in a riverine habitat in a rainforest. Some of the (undistributed) type material is heavily clothed in epiphytic mosses, Hymenophyllaceae, and lichens.
Conservation status. Eugenia malcomberi occurs in the Andohahela protected area (parcel I). No indication was provided on the label about its relative abundance at the time it was collected. Google™ Earth imagery (April 2013) shows a more or less continuous band of primary forest extending ca. 42 km south and ca. 135 km northeast, and ca. 10 km wide (east to west) at comparable elevations from the type locality. Considering its occurence in a nature reserve in fairly rugged terrain with considerable topographic relief, but also because it is known only from a single subpopulation with an AOO of 9 km 2 , Eugenia malcomberi is assigned a preliminary risk of extinction of "Vulnerable" [VU D2] following the IUCN Red List Categories and Criteria (IUCN 2012).
Comments. Th e petals during anthesis are quite large relative to the size of the fl ower buds, suggesting that they enlarge rapidly during early anthesis. Description. Trees to 8 m tall. Foliage glabrous and without evident oil glands except as noted. Branchlets round to somewhat laterally compressed, drying brown to light brown; bark smooth but cracking slightly longitudinally. Leaves stiffl y coriaceous, opposite or sometimes three per node, evenly distributed along branchlets, discolorous, somewhat glossy adaxially, matte abaxially. Axillary colleters absent. Petioles 25-35 mm, terete or slightly sulcate. Leaves 16-18 × 6.5-10.5 cm, elliptic, base slightly cuneate to mostly rounded, apex obtuse, surface and margin more or less fl at; adaxial surface with impressed midvein in lower ½ to 2/3, becoming fl ush distally; abaxial surface with dense but small oil glands (faint, use high magnifi cation); secondary veins indistinct, arising at 40-50° angle but mostly too faint to count with confi dence; intramarginal vein indistinct to barely visible, 1.5-2.5 mm from margin at midpoint of blade. Infl orescence up to 7 cm, structure uncertain but evidently of triads or botryoids, these solitary in leaf axils. Pedicels 4-14 mm, stiff and thick. Flowers mostly unknown. Calyx lobes (from dried fruit) 4, 3-5 mm, broadly rounded (much broader than long in fruit), persistent in fruit. Fruit 25-30 × 25-30 mm, globose.

Eugenia manomboensis
Etymology. Th e specifi c epithet is derived from the Réserve Spéciale de Manombo, the only known occurrence of this prominent species.
Phenology. Fruiting mid September; likely fl owering by August (unconfi rmed). Distribution. Known only from the type gathering in south-eastern Madagascar in Fianarantsoa from Réserve Spéciale de Manombo, near the coast (Fig. 7). Habitat and ecology. Dense, humid, low-altitude forest over lateritic soils. Conservation status. Eugenia manomboensis occurs in the Réserve Spéciale of Manombo. Considering its occurrence in a natural reserve, but also because it is known only from single subpopulation (AOO of 9 km2), Eugenia malcomberi is assigned a preliminary risk of extinction of "Vulnerable" [VU D2] following the IUCN Red List Categories and Criteria (IUCN 2012).
Comments. Large-leaved but indetermined speicmens of Malagasy Eugenia with large fruits have not been matched closely with fl owering material in some cases. However, given that none of the larger-leaved species have petioles as thick or as long as Eugenia manomboensis, no others are known from near the type locality, and no largeleaved species present globular fruits of this size, it appears that this species has not been named previously.
It seems likely that the large fruits of this tree are consumed by local widelife.
Etymology. In reference to the shape of the leaf blades. Vernacular name. Ropsay (Rabenantoandro 366). Phenology. Flowering September, November, and January; fruiting August through December.
Distribution. East-central Madagascar in Toamasina Province and south-eastern in Toliara Province from Mandena to Sainte Luce (Fig. 11).
Habitat and ecology. Littoral forest over sand; elevation ca. 5-53 m. Conservation status. With an EOO of 8,333 km 2 , an AOO of 63 km 2 and four subpopulations, none of which are situated within the protected network, Eugenia obovatifolia is assigned a preliminary risk of extinction of "Endangered" [EN 2ab(i, iii, iv)] following the IUCN Red List Categories and Criteria (IUCN 2012). Th e new species seems to be restricted to the highly threaten littoral forests and none of the known populations are currently protected.
Comments. Eugenia obovatifolia has a general resemblance to E. arenicola H. Perrier, E. hazompasika H. Perrier, and E. cloiselii H. Perrier given its ramifl orous, more or less fasciculate infl orescences. Th e four species also grow in relative proximity in southeastern and eastern Madagascar, often in sandy littoral forests, and may form a species complex that ranges approximately 1300 km between Taolognaro and Sainte Luce along the coast and somewhat inland to Vatomandry.
Eugenia obovatifolia diff ers from E. arenicola by its more or less glabrous leaves, whereas those of the latter are densely ferrugineous-hairy upon emergence and retain much of the indumentum on the abaxial surface. Eugenia obovatifolia diff ers each from E. cloiselii and E. hazompasika by virtue of the glabrous (or nearly so) infl orescences, which contrast with the hairy infl orescences of the latter two.
Etymology. From Ranomafana National Park. Phenology. Fruiting October and November; fl owering May. Distribution. Known from three collections in Ranomafana National Park in Fianarantsoa Province, southeast-central Madagascar (Fig. 4).
Conservation status. With only three collections known, an AOO of 9 km 2 and one subpopulation, which is situated within the protected area network (Ranomafana), Eugenia ranomafana is assigned a preliminary risk of extinction of "Vulnerable" [VU D2] following the IUCN Red List Categories and Criteria (IUCN 2012).
Comments. Th is species was fi rst recognized as undescribed by Turk (1997) as part of a dissertation studying plants in Ranomafana National Park, who approved of and collaborated with its publication here.
Phenology. Flowering January through April; fruiting March through November. Distribution. In the northern mountains of Madagascar in Antsiranana and Mahajanga provinces (Fig. 4).
Habitat and ecology. In mid-elevation, wet eastern forests often over granite or gneiss; elevation approximately 630-1880 m but needing confi rmation.
Etymology. It is a pleasure to name this species in honor of Désiré Ravelonarivo (b. 1966), a prolifi c collector of specimens in Madagascar. Désiré is native to the An-

Conservation status.
With an EOO of 436 km 2 , an AOO of 36 km 2 and three subpopulations, two of which are situated within the protected area network (Anjanaharibe-Sud), Eugenia ravelonarivoi is assigned a preliminary risk of extinction of "Vulnerable" [VU D2] following the IUCN Red List Categories and Criteria (IUCN 2012).
Comments. Th e label of the type gathering indicates "rougeâtres" (reddish) for petals, but this is doubtful and probably was meant for pinkish, which accords with information on the label of a paratype.
Among existing species, the leaf morphology of Eugenia ravelonarivoi resembles that of E. alatroensis H. Perrier and E. vatomandrensis H. Perrier, but the bases of their leaves are subsessile to sessile, which contrasts with the distinctly petiolate leaves of E. ravelonarivoi. In addition, the often cuspidate-caudate aspect of the leaf apex of new species is much more pronounced than of those two species, whose tips are barely (if at all) cuspidate.
Eugenia antongilensis H. Perrier has cuspidate apices on the blade, but it is a much longer leaf with shorter petioles. Eugenia musicola H. Perrier also resembles E. ravelonarivoi, especially given the membranous to weakly coriaceous texture of the leaf blades; however, the pedicels of the former are much thinner and more lax, and its petioles are shorter and much less (if at all) sulcate adaxially. Eugenia diospyroides diff ers with its generally larger fl owers and leaves, and leaves having a rounded apex.
Th e new species also closely resembles Eugenia radicifl ora in leaf morphology, but that species is said to have solitary fl owers with pedicels only 2-3 mm long, which does not match the fasciculate brachyblasts of E. ravelonarivoi. Th e leaves of Eugenia ravelonarivoi also superfi cially resemble those of E. gandhii, but the latter has densely punctate leaf blades and solitary fl owers. Finally, Eugenia ravelonarivoi resembles closely some evidently undescribed taxa Dr. David Gordon, an entomologist at Pittsburg State University, indicates that the visible damage on the leaves of some specimens may be from leaf cutter bees (Family Megachilidae), which often damage leaves in this manner to obtain material for the construction of their nests. Description. Trees to 10 m; dbh up to 13 cm; bark of main bole unknown. Plants glabrous throughout. Branchlets laterally compressed; youngest epidermis drying light brown to light gray but soon peeling and fl aking to reveal reddish-brown color (dried); oil glands of epidermis prominent and common. Leaves stiffl y coriaceous, mostly concentrated near branch tips, medium green and slightly discolorous, irregularly sinuous, surfaces matte. Axillary colleters absent. Petioles 3-5 mm, becoming slightly sulcate distally and broadening towards apex. Leaf blades 3.5-9.0 × 2.5-5.0 cm, broadly elliptic to obovate or broadly obovate, base cuneate, apex obtuse to somewhat acute; adaxial surface with midvein broadly but shallowly sulcate in the lower half, becoming fl ush distally; oil glands absent; abaxial surface oil glands common to dense but small and faint and fading with maturity (visible only with magnifi cation), secondary veins somewhat indistinct, intramarginal vein indistinct and ca. 0.5-1.0 mm from margin at midpoint of blade. Infl orescence terminal or axillary, monads 1-3 in each axil. Pedicels 15-25 mm, relatively thick (1.2-2.0 mm at maturity), ascending to erect, rigid. Bracteoles 2, 1-2 mm, ovate, stiff , persistent and more or less divergent in fruit. Hypanthium 3.5-5 mm, cupulate, oil glands common to dense. Calyx lobes 4, 5-8 mm, broadly rounded, apex obtuse, densely glandular, persistent and crowning the fruit, stramineous or rufous-beige (fresh). Flowers unknown. Berries 1.5-1.8 × 2.0-2.3 cm, subglobular to globular, densely glandular, slightly verrucose (dried), yellowish (possibly not fully mature).
Etymology. It is a pleasure to name this species in honor of Richard Razakamalala (b. 1962), a Malagasy botanist who has collected over seven thousand specimens of plants from Madagascar since 2001. His prolifi c collecting eff orts have contributed signifi cantly to our increased knowledge of the Malagasy fl ora. Four of his Eugenia collections are designated as type specimens in this publication.
Phenology. Fruiting late February. Distribution. Known only from the type gathering in Antsiranana Province, north-eastern Madagascar, about 14 km west of the coast in the Masoala Peninsula (Fig. 2).
Habitat and ecology. In dense humid lowland forests over quartzite from ca. 35 m elevation.

Conservation status.
With only one collection known from just outside the Masoala protected area, Eugenia razakamalalae is assigned a preliminary risk of extinction of "Endangered" [EN 2ab(i,iii,iv)] following the IUCN Red List Categories and Criteria (IUCN 2012). It is highly probable that the new species occurs within Masoala National Park, and is therefore protected.
Comments. Th e leaf morphology of Eugenia razakamalalae resembles that of Eugenia arthroopoda H. Perrier var. ambalavensis H. Perrier, an invalid name given the lack of a Latin diagnosis in the protologue (Perrier de la Bâthie 1953a). However, the pedicels, hypanthium and calyx lobes of Humbert 22,804 (a syntype of var. ambalavensis) have a dense, uniform, appressed rusty indumenta that is also present on the outer surface of the petals, apart from an approximately 1 mm wide glabrous margin. In contrast, the pedicels in fruit of E. razakamalalae are glabrous (fl owering material is unknown).
Distribution. Known from the north-eastern coastal region of Madagascar in Antsiranana Province, from Orangéa to about 140 km along the coast near Vohemar (Fig. 2).
Habitat and ecology. Littoral forests over sand, inland from the ocean shore less than 1 km; elevation 10 m.
Conservation status. With an EOO of 358 km 2 , an AOO of 36 km 2 and three subpopulations, one of which is situated within a proposed protected area that currently holds a temporary protection status (Oronjia), Eugenia richardii is assigned a preliminary risk of extinction of "Endangered" [EN B1ab(iii)+ B2ab(iii)] following the IUCN Red List Categories and Criteria (IUCN 2012). Th e species seems to have a highly restricted distribution in dry deciduous forests over sand. Th e area around Oronjia is mostly degraded as a result of human activities and is under serious threat. A defi nitive protection of the dry deciduous forests over sand of Oronjia would likely enable the species to be downlisted from the current risk of extinction to "Vulnerable".
Comments. Eugenia richardii most closely resembles two species, including Eugenia vanwykiana N. Snow, which diff ers by its larger leaves with cuneate bases (Snow et al. 2012). Th e second fairly close match is Eugenia calciscopulorum, which occurs signifi cantly farther inland (ca. 11 km) than E. richardii and at a higher elevation (ca. 410 m) and over limestone, compared to the sandy substrates of E. richardii. Th e leaves of E. calciscopulorum have more pronounced secondary veins above and below, retain the pronounced glands into the mature leaf stage, and have a more pronounced petiole.
Eugenia richardii also resembles E. quadrifl ora H. Perrier, which is known from a single collection in 1961 from east of Lake Alaotra. Both species have thickly coriaceous, densely glandular leaf blades with irregularly undulating surfaces and moderately to strongly recurved margins, and simple fl owers arising terminally. However, the leaves of E. quadrifl ora are sessile with strongly and irregularly recurved blades (although this may refl ect to some degree its condition when pressed and dried); in contrast, the leaf blades of of E. richardii are distinctly but shortly petiolate and slightly recurved.
Elsewhere, the leaf morphology of Eugenia richardii somewhat resembles that of Eugenia neofasciculata Bennet from Mauritius (formerly E. fasciculata Guého & A.J. Scott). Based on comparison digitally with an isotype (Lorence DL 1214 (P [P00390098]!), the species are similar by virtue of stiffl y coriaceous, densely punctate, obovate-elliptic leaf blades with slightly undulating surfaces and recurved margins that arise from relatively short petioles. However, whereas the adxial petiolar surface of E. richardii is fl at, that of E. neofasciculata is somewhat sulcate. In addition, the infl orescence of E. richardii is a terminal pair of fl owers with one arising in each leaf axil, whereas that of E. neofasciculata is conspicuously fasciculate (mostly) on naked branches or within leaf axils.
Etymology. Th e specifi c epithet is derived from elements of the native language of northern Madagascar, in which ampoka means mosses. Eugenia tiampoka thus means the "Eugenia that likes mosses".
Phenology. Flowering October through February; fruiting by early April, likely commencing by March and extending through May.
Distribution. Northern Madagascar in Antsiranana Province in the northern mountains (Fig. 2).
Habitat and ecology. Mid-to high-altitude humid forests laden with mosses, sometimes along river banks; 1990-2300 m in the northern highlands of Madagascar. A team of botanists lead by MC conducted a series of fi eld expeditions between 2005 and 2008 to explore the fl ora and vegetation of this long-neglected region, during which more than 4,400 collections were made, yielding material of several plant species new to science (Callmander et al. 2008(Callmander et al. , 2009(Callmander et al. , 2012. Conservation status. With an EOO of 71 km 2 , an AOO of 45 km 2 and two subpopulations, neither of which is within the protected area network, Eugenia tiampoka is assigned a preliminary risk of extinction of "Endangered" [EN B1ab(iii)+2ab(iii)] following the IUCN Red List Categories and Criteria (IUCN 2012).
Comments. Th e leaves reportedly are aromatic (Birkinshaw 950, Rakotovao 2398). One specimen is said to have a nectariferous disk (Randriarivelo 350). A specimen of 30 m stature (Ravelonarivo 647) from the same general area and also said to be aromatic, but growing at 1700 m, has a slightly diff erent aspect to the leaves, also may be this species.
Th e morphological gestalt of the leaves and infl orescences resemble most closely those of E. thouvetiana H. Perrier, which however lacks the punctate oil glands of the abaxial leaf surface, and which has much longer pedicels subtending the individual fl owers than those of E. tiampoka. Th e type locality of E. thouvetiana is located in Analamazaotra-Andasibe in Toamasina Province, some 525 km south of the southernmost known collection of E. tiampoka. Eugenia tiampoka also somewhat resembles Eugenia hazompasika H. Perrier, the latter of which has much longer, broader and stiff er leaves with a less abruptly cuneate base.
Etymology. In honor of Dr. Peter G. Wilson (b. 1950) of the Royal Botanic Gardens in Sydney, Australia; colleague, collaborator, and long-time student of Myrtaceae (e.g., Wilson and Waterhouse 1982;Wilson et al. 2005;Snow and Wilson 2010;Wilson 2011 Distribution. East-central Madagascar in Toamasina Province, occurring near Analamazaotra National Park (Fig. 2).
Habitat and ecology. In humid, sometimes disturbed forests at middle altitudes from ca. 980-1103 m.
Conservation status. With fi ve collections known, an AOO of 9 km 2 and one subpopulation that is situated outside the protected area, Eugenia wilsoniana is assigned a preliminary risk of extinction of "Critically Endangered" [CR A3c+B1ab(iii)] following the IUCN Red List Categories and Criteria (IUCN 2012). Current data suggest the species has a narrow distribution, and the montane evergreen tropical forests where the species grows are threatened by mining activities. Th e discovery of this species in one of the nearby protected areas probably would allow downlisting to "Endangered".
Comments. Th e leaf and fl oral morphologies of Eugenia wilsoniana are suggestive of Eugenia diospyroides H. Perrier. However, E. wilsoniana has much shorter leaves and a hairy staminal ring, in contrast with the much larger leaves and glabrous staminal ring of E. diospyroides (e.g., Randriatifi ka 118 et al. [KSP]). Th is new species resembles even more closely another newly described species herein, E. bemangidiensis, which differs by its cauliforous infl orences occurring in tight, relatively short fascicles, secondary veins on the leaf blades arising at steeper angles, and occurring in a diff erent habitat at much lower elevations some 750 km southeast from the presently known occurrences of E. wilsoniana. Moreover, the foliage and fl oral parts of E. bemangidiensis are entirely glabrous, the abaxial laminar midvein protrudes less prominently, the adaxial laminar sulcus is not as deep or narrow, and its adaxial petiolar sulcus is broader and shallower than those of E. wilsoniana.

Lectotypifications
Many species of Eugenia in Madagascar described by Perrier de la Bâthie (1953a) eventually will require lectotypifi cation to establish coherent species concepts and avoid nomenclatural confusion. We have reviewed all of the existing published small-leaved Eugenia species in Madagascar and the currently available specimens, and have seen some additional material that has helped to clarify species delimitations. Here we lectotypify three names to help clarify species' boundaries and nomenclature.
Lectotypifi cation is particularly important for Eugenia scottii, since the protologue included twelve syntype collections that clearly included more than one taxon. Eugenia hovarum H. Perrier and Eugenia scottii H. Perrier were described for relatively smallleaved species occuring mostly in the mountainous parts of Madagascar, and both were based on multiple syntypes (Perrier de la Bâthie 1953a). Perrier de la Bâthie (1953a: 169) referred to Eugenia scottii as having a large number of forms that he could not distinguish taxonomically ("…un très grand nombre de formes que nous n'avons pu distinguer"). In light of the material now available, we believe that the specimens can be assigned comfortably to fi ve diff erent species, four of which already have valid names. Among the syntypes, we have chosen Humbert 18501 at 2300 m elevation from the Tsaratanana Massif in Antsiranana Province as the lectotype because it is among the better fl owering syntype collections, and represents a species that cannot be assigned to any other published taxon. None of the syntypes at isolectotype: P [P00118031]!). Syntype: MADAGASCAR. Prov. Antananarivo: Imerina, comm. 20 Nov. 1889, Campenon s.n. (P [P00118033]!).

Eugenia scottii
Perrier de la Bâthie (1953a: 170) only cited two syntypes that clearly belong to species that we consider to be Eugenia hovarum. Not atypically, Perrier de la Bâthie made no indication of which should be regarded as the holotype, but his own collection (Perrier de la Bâthie 14007) is the more complete of the two, comprises two separate sheets at P, and bears a detailed collection locality information, unlike the other syntype, and is here designated as the lectotype.
Additional Th e species concept for Eugenia nompa H. Perrier has been uncertain. Because the syntypes were clearly designated by Perrier de la Bâthie (1953a), we lectotypify one of these to clarify its diagnostic characters, which include: leaf blades narrowly elliptic, apex acute; midvein deeply and narrowly sulcate lower 75%+; margins somewhat sinuous; infl orescence of somewhat elongate cymes or sparsely-fl owered botryoids, terminal or axillary in clusters, the whole (at least when young) covered with a ferrugineous indumentum.
Perrier de la Bâthie (1953a) also described Eugenia nompa var. arborea H. Perrier, an invalid name based on Perrier de la Bâthie 16261. Th e infl orescence structure of those specimens diff er considerably from the lectotype recognized above, and it is still unclear whether specimens designated as this variety can be confi dently placed in extant taxa.

Discussion
Th e eighteen species of Eugenia newly proposed here increases the known number of native species in Madagascar from approximately 46 to 68, all of which are endemic. Of these, 32 (approximately half ) have been described in the last fi fteen years (Miller 2000;Snow 2008Snow , 2011Snow et al. 2012 and this paper). Specimens of the introduced and possibly naturalized Neotropical species Eugenia unifl ora L. and E. brasiliensis Lam. have been confi rmed by the fi rst author (housed at MO). Both species produce edible fruits, and the former also is used as a hedgerow. Th e paucity of collections suggests that these species are not widely planted.
Many of the original descriptions by Perrier de la Bâthie (1953a) of Malagasy Eugenia were relatively incomplete, and many of his infraspecifi c names are invalid. Because preliminary studies suggest some of those infraspecifi c taxa are worthy of recognition, new names will be necessary in future combinations after additional study.
Malagasy Eugenia have a relative paucity of easily diagnostic, qualitatively diff erent morphological character states compared to some baccate genera of Myrtaceae (e.g., Snow et al. 2003;Snow 2008), including the related genus Syzygium. For example, few species of Eugenia have winged or quadrangular branchlets, apically attenuate calyx lobes, costate fruits, or other qualitatively distinct character states that recur in many other baccate genera, and which simplify identifi cations and the writing of keys (e.g., Snow 2008;Snow et al. 2001Snow et al. , 2008. Although additional undescribed species are known with a high degree of confi dence, further research and more collections will be needed to produce a complete taxonomic treatment of the genus for Madagascar.
Despite additional new species of Eugenia remaining to be described and other gaps in knowledge of Malagasy taxa, tentative patterns of distribution, ecological preferences and evolution are beginning to emerge . One: Moderately high levels of specifi c diversity of Eugenia can occur in relatively localized areas, such as the approximately 15 species (including some undescribed) between Taolagnaro and Sainte Luce on the southeast coast. Two: Most Eugenia occupy one of three general areas: i) high altitudes in the northern mountainous massifs; ii) humid forests in the vicinity of eastern escarpment; or iii) in drier lowland habitats near the eastern coast. Th ree: Most species of Eugenia in Madagascar have relatively limited geographic ranges. Th is contrasts with some Malagasy species of Syzygium Gaertn., the other genus of Myrtaceae native to Madagascar, which includes some widely distributed species, such as S. bernieri  (Byng 2014;Byng et al. in press). Given that many species of Eugenia are relatively short trees or shrubs, in contrast with many of Syzygium that are canopy or emergent species, the average dispersal distance of fruits of Eugenia species may be considerably less than Syzygium. Four: No species of Eugenia is an ecological dominant, although some species can be locally common, such as E. echinulata and E. obovatifolia. Five: Caulifl ory (fl owers arising on main bole) and ramifl ory (fl owers arising on naked branches below leaves) are common methods of fl oral presentation, and often occur largely in lieu of terminal or axillary presentations. Caulifl ory and ramifl ory presumably refl ect selective pressures for pollination and/or fruit dispersal. Six: All known species have whitish, cream-colored, or pinkish-magenta petals. In contrast, red, yellow, orange, purple or blue petals are unknown. While almost nothing is known about the pollination biology of Malagasy Eugenia, the petal colors likely refl ect selection pressures of generalized pollinators such as bees. Seven: Selection has sometimes favored relatively large fruits and embryos (e.g., E. andapae, E. manomboensis, E. iantarensis, E. guillotii, E. manomboensis, E. ravelonarivoi), which suggests consumption of and dispersal by mammals or large birds. Lemurs (this paper; Snow et al. 2012) are among the known consumers, and information concerning bird consumption would be appreciated by the fi rst author. Field studies documenting the pollinators of fl owers and consumers of fruits of Eugenia (and Syzygium) would be valuable graduate theses. Eight: Although dioecy in Eugenia is well known from western and southern Africa and adjacent areas (van Wyk and Lowry 1988;Bernardini et al. 2014, J. Byng, pers. comm. 2014, this reproductive system has not yet been documented among Madagascar members of Eugenia. Many species of Malagasy Eugenia, including several proposed here, are Th reatened, Endangered, or Critically Endangered following IUCN Criteria and Recommendations (IUCN 2012). In view of rapidly continuing deforestation in Madagascar, the occurrence of many from fi ve or fewer collections, and that some species are known from only fl owering or fruiting material, additional collecting of Eugenia should remain a high priority.
We believe that at least ten or more additional species remain to be described, and stress here that more material of other species would enhance our knowledge of taxonomy, distribution, and conservation priorities. Duplicate material of Malagasy Eugenia sent to the fi rst author at the T.M. Sperry Herbarium (KSP) for future studies would be greatly appreciated.
Corrigendum: Th e correct collection number for the paratype specimen of Eugenia vanwykiana N. Snow is Razakamalala 1234 et al. Th e protologue incorrectly cited the collection number as 1324 (Snow et al. 2012).